PV Cruz, EL Belmont, R Boldrini, N Hamada

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Neotropical Entomology

journal homepage: www.scielo.br/ne ISSN: 1519-566X

SyStEmatIcS, morphology aNd phySIology

leptohyphidae (Insecta: Ephemeroptera) from Northeastern Brazil

PV Cruz, EL Belmont, R Boldrini, N Hamada

Lab de Citotaxonomia e Insetos Aquáticos, Instituto Nacional de Pesquisas da Amazônia – INPA, 69011-970 Manaus, AM, Brasil

Keywords

Aquatic insect, Ephemerelloidea, Tricorythopsis, Tricorythodes

Correspondence

Paulo V Cruz, programa de pós-graduação em Entomologia (PPGENT), Instituto Nacional de Pesquisas da Amazônia – INPA, Av. André Araújo 2936, Bairro Petrópolis, 69011-970 Manaus, AM, Brasil; pvilelacruz@gmail.com Edited by Roberto A Zucchi – ESALQ/USP Received 07 March 2011 and accepted 26 July 2011

Abstract

New records of Leptohyphidae for Northeastern Brazil are provided. Previously, only Tricorythopsis bahiensis Dias, Salles & Ferreira had

been recorded in this region, but we now record one species of

Amanahyphes Salles & Molineri, one of Leptohyphes Eaton, three of Traverhyphes Molineri, three of Tricorythodes Ulmer, and seven of Tricorythopsis Traver. Two of these species are recorded for the first

time in Brazil. As all of these species are reported, the Northeastern Brazil became the second most diverse geographic region in Brazil in Leptohyphidae.

Introduction

Leptohyphidae is considered to be one of the most diverse families of Ephemeroptera in South America, after Baetidae, Leptophlebiidae and Polymitarcyidae (Salles 2006). Leptohyphidae has a Pan-American distribution and is represented by approximately 138 species (Barber-James et al 2008), 35 of which, belonging to Amanahyphes, Leptohyphes, Leptohyphodes, Macunahyphes, Traverhyphes, Tricorythodes and Tricorythopsis, are reported for Brazil

(Salles et al 2010a, b).

There is a lack of knowledge of the Ephemeroptera fauna in the Center-West and Northeastern Brazil, especially for the family Leptohyphidae (Salles et al 2004).

The Northeast is one of the least sampled regions of Brazil, and consequently it has the most incipient taxonomic knowledge on Ephemeroptera when compared to the Southern and Southeastern Brazil (Lima et al 2010). In

most cases, species distribution can only be accessed in often old and not easily accessible descriptive studies (e.g. Allen 1967, 1977). The lack of information of this

nature limits the analysis of biogeographic patterns and

precludes the development of regional identification keys

and systematic studies. Therefore, our objective is to contribute to the increasing knowledge of the distribution of the Leptohyphidae fauna in Northeastern Brazil.

Material and Methods

Collections were done between July and August 2010 in four northeastern states of Brazil: Maranhão, Piauí, Ceará and Bahia. Additionally, samples collected in the state of Goiás, Center-West of Brazil, were included because the sampled area is located near the border with state of Bahia (Fig 1). Data pertaining to each sampled locality are provided in Table 1.

Nymphs were collected with an aquatic entomological net, while adults were captured with light traps.

Specimens were fixed in 80% ethanol and permanent

slides, whenever needed, were prepared in Euparal®.

Species identification was based on Domínguez et al

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(M165C) stereomicroscope with a DFC420 digital camera.

Species that had only one specimen collected were deposited in the Coleção de Invertebrados do Instituto

Nacional de Pesquisas da Amazônia – INPA, Manaus, Amazonas, Brazil. Species represented by several specimens also had specimens deposited in the Coleção de Invertebrados do Centro Universitário Norte do Espírito

Fig 1Map of Northeastern states and Goiás showing the sampled sites. (BA = Bahia, CE = Ceará, GO = Goiás, MA = Maranhão, PI = Piauí)

Table 1 Sampled sites in Northeastern region of Brazil and in Goiás state near the border with Bahia state, followed by locality, geographic coordinates, date and collectors.

Sample suite State locality geographic coordinates date collectors

pt 01 maranhão ribamar riquente, ma-010, rio lajeado 06° 04’ 15.6’’S 47° 22’

56.6’’W 21.vii.2010 Nh, pVc, rB

pt 02 maranhão porto Franco, ma-010, rio Farinhas 06° 31’ 47.3’’S 47° 28’

11.4’’W 22.vii.2010 Nh, pVc, rB

pt 03 maranhão carolina, disable hydroelectric, rio Itapecuru

07º 24’ 53.2’’S 47º 12’

54.9’’W 22.vii.2010 Nh, pVc, rB

pt 04 piauí São João da Fronteira, rio Jenipapo 04° 00’ 15.8’’S 41° 26’

53.8’’W 26.vii.2010 Nh, pVc, rB, rQ

pt 05 ceará Ubajara, rio Jaburu, Boi morto Waterfall 03° 52’ 36.2’’S 41° 10’

0.08’’W 26.vii.2010 Nh, pVc, rB, rQ

pt 06 Bahia camacan, reserva particular de patrimônio Natural Serra Bonita

15° 22’ 59.1’’S 39° 33’

21.2’’W 02.viii.2010 Nh, rB, rQ

pt 07 Bahia rio de contas, pico do Itobira, rio comburú 13° 24’ 19.1’’S 41° 52’

53.9’’W 04.viii.2010 Nh, rB, rQ

pt 08 Bahia piatã, rio de contas 13° 07’ 23.6’’S 41° 50’

14.1’’W 05.viii.2010 Nh, rB, rQ

pt 09 Bahia correntina, rio correntina, near the restaurant

13° 19’ 59.6’’S 44° 36’

08.0’’W 07.viii.2010 Nh, rB, rQ

pt 10 Bahia correntina, rio correntina, near posto cachoeira

13° 31’ 09.6’’S 45° 21’

01.8’’W 08.viii.2010 Nh, rB, rQ

pt 11 goiás posse, rio do prata, Bahia/goiás border 14° 14’ 46.4’’S 46° 33’

07.0’’W 09.viii.2010 Nh, rB, rQ

pt 12 Bahia correntina, prainha community, rio arrojado

13º 31’ 25.1’’S 44º 43’

30.7’’W 05.viii.2003 Nh, FFS 1029 km

N

go

Ba ma

pt1 pt2 pt3

pt5 pt4

pt11PT10 pt12 pt9 PT7pt8

pt6

cE

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Fig 2 Dorsal habits of nymphs: a) Amanahyphes saguassu; b) Leptohyphes petersi; c) Traverhyphes

(Mocohyphes) edmundsi; d) Traverhyphes

(M o c o h y p h e s) y u a t i; e ) Tr a v e r h y p h e s

(Traverhyphes) indicator; f ) Tricorythodes mirca; g) Tricorythodes quizeri; h) Tricorythopsis araponga; i) Tricorythopsis minimus; j)

Tricorythopsis pseudogibbus; k) Tricorythopsis chiriguano.

Santo – CEUNES, São Mateus, Espírito Santo, Brazil. For each species recorded, data on the known stages, distribution with emphasis on Brazil, comments (if necessary), and a list of examined material is provided. Abbreviations and symbols used: “I” = imago, “N” = nymph, “PT” = collection site.

Results

A total of 159 specimens belonging to 15 species in five

genera were collected.

amanahyphes Salles & Molineri

Amanahyphes saguassu Salles & Molineri (2006) (Fig 2a) Known stages.I♂, N

Distribution. Brazil: states of Amazonas (Manaus and

Presidente Figueiredo municipalities) (Salles & Molineri 2006), Pará (Parauapebas municipality) (Gonçalves &

Da-Silva 2010); new record:_{state of Maranhão (Carolina}

municipality).

Comments. This is a very common species in the states

of Amazonas and Pará, usually collected on roots with sediment.

Material examined. PT3: (1N), 22.vii.2010, INPA.

leptohyphes Eaton

Leptohyphes petersi Allen (1967) (Fig 2b)

Known stages.I♀♂, N.

Distribution. Peru (Allen 1967); new record_{: Brazil:}_state

of Bahia (Camacan municipality).

Material examined. PT6: (4N), 02.viii.2010, two in INPA, two in CEUNES.

Traverhyphes Molineri

Traverhyphes (Mocohyphes) edmundsi (Allen 1973) (Fig 2c)

a

e

b

f

i

c

g

j

d

h

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Known stages.I♀♂, N.

Distribution. Argentina (Molineri 2004); Brazil: states of

Paraná (Morretes, Curitiba, Sete Quedas, Prudentópolis and Guairá municipalities) (Allen 1973, Dias et al 2007a),

São Paulo (Intervales, Campos do Jordão and Salesópolis municipalities) (Molineri 2004, Dias et al 2007b), Rio

Grande do Sul (Cachoeira do Sul municipality) (Allen 1973, Dias et al 2007a), Santa Catarina (Joinville and Rio

Warnow municipalities) (Allen 1973, Dias et al 2007a);

new records:_{states of Maranhão (Ribamar Riquente}

and Porto Franco municipalities), Bahia (Camacam municipality), and Goiás (Posse municipality).

Material examined. PT1: (5N), 21.vii.2010; PT2: (3N),

22.vii.2010; PT6: (5N), 02.viii.2010; PT11: (1N), 09.viii.2010, seven in INPA, seven in CEUNES.

Traverhyphes (Mocohyphes) yuati Molineri (2004) (Fig

2d)

Known stages.N.

Distribution.Argentina (Molineri 2004);Brazil: states of

São Paulo (Campos do Jordão, Salesópolis and Iporanga municipalities) (Molineri 2004, Dias et al 2007b), Rio

de Janeiro (Nova Friburgo and Itatiaia municipalities) (Molineri 2004, Dias et al 2007a), Minas Gerais and

Espírito Santo (Espera Feliz municipality) (Salles et al

2010a); new records: states of Maranhão (Ribamar

Riquente and Porto Franco municipalities), Piauí (São João da Fronteira municipality) and Goiás (Posse municipality).

Comments. The collected nymphs have a series of

characteristics that deviate from the original description of T. yuati. However, according to Carlos Molineri

(personal communication), nymphs of this species can have large morphological variation. The differences observed were: 1) coxae with black mark; 2) femur with subapical black mark dorsally and on outer and inner surface; 3) maxillary palp 2-segmented, very long, more

than 2.5× longer than wide, with fine and long apical seta;

4) tarsal claws with 5-7 marginal denticles and 4+3-2 submarginal denticles, with apical seta.

Material examined. PT1: (10N), 21.vii.2010; PT2:

(5N), 22.vii.2010; PT4: (1N), 26.vii.2010; PT11: (1N), 09.viii.2010, nine in INPA, eight in CEUNES.

Traverhyphes (Traverhyphes) indicator (Needham &

Murphy 1924) (Fig 2e)

Known stages. I♀♂, N.

Distribution. Argentina (Needham & Murphy 1924,

Molineri 2001a), Uruguay (Traver 1958), Brazil: states of São Paulo (Ribeirão Preto, Ribeirão Grande and Iporanga municipalities) (Dias et al 2007a, b) and Espírito

Santo (Pinheiros municipality) (Salles et al 2010a);

new records: states of Maranhão (Ribamar Riquente

and Porto Franco municipalities), Bahia (Correntina municipality) and Goiás (Posse municipality).

Material examined. PT1: (8N), 21.vii.2010; PT2: (2N),

22.vii.2010; PT9: (4N), 07.viii.2010; PT10: (2N), 08.viii.2010; PT11: (1N), 09.viii.2010, seven in INPA, ten in CEUNES.

Traverhyphes (Traverhyphes) sp.1

Known stages.I♂.

Distribution.Brazil: new record:_{state of Ceará (Ubajara}

municipality).

Comments.Only one male imago was collected in a light trap. The specimen was in poor condition,preventing us from identifying the species.

Material examined.PT5: (1I), 26.vii.2010, INPA.

tricorythodes Ulmer

Tricorythodes mirca Molineri (2002) (Fig 2f)

Known stages. I♀♂, N.

Distribution. Bolivia (Molineri 2002); Brazil: states of Espírito Santo (São Mateus and Sooretama municipalities) (Salles et al 2010a); new records:_{states of Maranhão}

(Ribamar Riquente and Porto Franco municipalities) and Piauí (São João da Fronteira municipality).

Comments. Tricorythodes mirca and T. arequita Traver have similar nymphs. Tricorythodes mirca has tarsi without black mark, while the nymph of T. arequita has tarsi with black mark. However, some of our T. mirca

specimens have a small black mark on the tarsi. Other variations observed in the specimens collected in this study were: 1) tarsal claws with 8–11 marginal denticles, without submarginal denticles (as previously cited by Salles et al 2010a); 2) maxillary palp with apical and subapical setae.

Material examined. PT1: (2N), 21.vii.2010; PT2: (21N), 22.vii.2010; PT4: (6N), 26.vii.2010, 14 in INPA, 15 in CEUNES.

Tricorythodes quizeri Molineri (2002) (Fig 2g) Known stages. I♀♂, N.

Distribution. Bolivia (Molineri 2002); Brazil: state of Mato

Grosso (Nova Xavantina municipality) (Dias et al 2009);

new record: state of Goiás state (Posse municipality).

Comments. Since the locality where the specimens were

collected is near to the border with Bahia, is likely that this species may also occurs in the latter state.

Material examined. PT11 (1N), 09.viii.2010, INPA.

Tricorythodes sp. 1

Known stages.N.

Distribution. new records:_{Brazil: states of Bahia}

(Correntina municipality) and Maranhão (Ribamar Riquente and Porto Franco municipalities).

Comments. It was not possible to identify the collected specimens. They probably represent an undescribed species.

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22.vii.2010; PT9 (1N), 07.viii.2010, three in INPA, three in CEUNES.

tricorythopsis Traver

Tricorythopsis araponga Dias & Salles (2005) (Fig 2h) Known stages.N.

Distribution.Brazil: states of Espírito Santo (Jerônimo

Monteiro municipality) (Dias & Salles 2005), Minas Gerais (Araponga municipality) (Dias & Salles 2005), Rio de Janeiro (Itatiaia municipality) (Dias & Salles 2005), and São Paulo (Salesópolis municipality) (Dias & Salles 2005); new record: state of Bahia (Rio de Contas

municipality).

Comments. The collected specimen has all diagnostic

characteristics of T. araponga, except tarsal claw with

6-8 marginal denticles and 5-8+8-10 submarginal denticles.

Material examined.PT7 (1N), 04.viii.2010, INPA.

Tricorythopsis bahiensis Dias, Salles & Ferreira (2008) Known stages.N.

Distribution.Brazil: states of Roraima (Arraiá municipality)

(Dias et al 2008), Amazonas (Presidente Figueiredo

municipality) (Dias et al 2008), and Bahia (Correntina

municipality) (Dias et al 2008); new record: state of

Maranhão (Ribamar Riquente municipality).

Materialexamined. PT1 (4N), 21.vii.2010; PT12 (7N),

05.viii.2003, five in INPA, six in CEUNES.

Tricorythopsis minimus (Allen 1973) (Fig 2i) Knownstages. I♀♂, N.

Distribution. Argentina (Molineri 2001b), Uruguay (Traver

1958), Brazil: states of Rio Grande do Sul (Panambi municipality) (Allen 1973) and Espírito Santo (Sooretama municipality) (Salles etal 2010a); new record: state of

Bahia (Correntina municipality).

Materialexamined. PT9 (1N), 07.viii.2010, INPA.

Tricorythopsis pseudogibbus Dias & Salles (2005) (Fig 2j) Knownstages.N.

Distribution. states of Rio de Janeiro (Itatiaia municipality)

(Dias & Salles 2005, Salles et al 2010a), Minas Gerais

(Araponga municipality) (Dias & Salles 2005, Salles et al

2010a), and São Paulo (unknown municipality) (Mariano & Polegatto 2011); new record: state of Bahia (Rio de

Contas municipality).

Materialexamined. PT7 (1N), 04.viii.2010, INPA.

Tricorythopsis chiriguano Molineri (2001a) (Fig 2k) Knownstages. I♀♂, N.

Distribution. Bolivia (Molineri 2001a); new record:

Brazil: state of Maranhão (Ribamar Riquente and Porto Franco municipalities).

Comments. Tricorythopsis chiriguano and T. minimus are

very similar; the abdominal color pattern and length of

operculate gills can be used to distinguish them.

22.vii.2010, two in INPA, three in CEUNES.

Tricorythopsis sp. 1

Knownstages.N.

Distribution. new record: _{Brazil: states of Maranhão}

(Ribamar Riquente municipality).

Comments. It was not possible to identify the collected specimens. They probably represent an undescribed species.

Materialexamined. PT1 (30N), 21.viii.2010, 15 in INPA, 15 in CEUNES.

Tricorythopsis sp. 2 Knownstages.N.

Distribution. new records:_{Brazil: states of Maranhão}

(Ribamar Riquente and Porto Franco municipalities) and Bahia (Correntina municipality).

Comments. It was not possible to identify the collected

specimens. They probably represent an undescribed species.

22.vii.2010; PT9 (1N), 07.viii.2010, seven in INPA, eight in CEUNES.

Discussion

The Northeast of Brazil represents nearly 18% of the

country area and it is the region that has the largest number of biomes: Amazon, Cerrado, Caatinga and Atlantic forest (IBGE 2004). Four types of climate can be found in this unique Brazilian region: equatorial humid, coastal humid, tropical and semi-arid (IBGE 2002).

Despite having a high diversity of biomes and climates, Caatinga, with a semi-arid climate and seven to 11 months of water shortage, is the predominant biome (Leal et al

2005). The available water resources are heavily used by the population of the region (Rebouças 1997). Caatinga is the only biome entirely restricted to Brazil’s national territory. However, little attention has been given to the preservation of this region, and the contribution of this area to the country’s biodiversity is high and has probably been underestimated (Silva et al 2004).

Only one inventory of the Ephemeroptera fauna has been done in the Northeast of Brazil, focusing on Baetidae from the state of Bahia (Lima et al 2010). Leptohyphidae

was represented in this region only by T. bahiensis, which

was described from Bahia (Dias, Salles & Ferreira 2008). With the new records we provided, the Leptohyphidae fauna from the Northeast of Brazil became one of the most

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two genera to the Center-West of Brazil.

Of the seven known Leptohyphidae genera recorded in Brazil, Macunahyphes and Leptohyphodes were the only

ones missing from the Northeast. Two of the 15 species reported in this study represent new species records for Brazil: L. petersi, previously known only to Peru, and T. chiriguano only to Bolivia. One of the specimens collected

(one imago represented by Traverhyphes (Traverhyphes)

sp.1) could not be identified to the species level because

of the poor conditions of specimen, but its record is important to the knowledge of the distribution of the genus in Brazil.

Our study reinforces the need to increase the sampling of aquatic insects in Brazil. We were able to considerably increase the number of known species for the Northeastern region, although the occurrence of a small number of streams and rivers sparsely distributed

in our sampling area. Certainly, a larger collection effort,

including more detailed samples in all of the biomes in

the Northeast of Brazil would significantly increase our

knowledge on the diversity of Ephemeroptera.

Acknowledgments

We thank Dr Carlos Molineri and Lidianne Salvatierra

for all the help offered during the preparation of this

paper, Dr Victor Becker allowed collection in the Serra Bonita RPPN, state of Bahia, Dr Ranyse Querino helped

us during field work and Dr Philip M. Fearnside reviewed

the manuscript. Financial support for travel was provided by CNPq and MCT-INPA-PPI, and laboratory infrastructure was provided by the PRONEX-CNPq/FAPEAM project.

References

Allen RK (1967) New species of New World Leptohyphinae (Ephemeroptera: Tricorythidae). Can Entomol 99: 350-375.

Allen RK (1973) New species of Leptohyphidae Eaton. Pan-Pacific Entomol 49: 363-372.

Allen RK (1977) A new species of Tricorythodes with notes

(Ephemeroptera: Tricorythidae). J Kansas Entomol Soc 50: 431-435.

Barber-James HM, Gattolliat JL, Sartori M, Hubbard MD (2008)

Global diversity of Mayflies (Ephemeroptera, Insecta) in

freshwater. Hydrobiologia 595: 339-350.

Dias L G, Cabette H S F, Souza D P (2009) A new species of

Tricorythodes Ulmer, 1920 (Ephemeroptera: Leptohyphidae)

and first record of Tricorythodes quizeri Molineri, 2002 from

Brazil. Aquat Insects 31: 95-99.

Dias LG, Molineri C, Ferreira PSF (2007a) Ephemerelloidea (Insecta: Ephemeroptera) do Brasil. Pap Avulsos Zool 47: 213-244.

Dias LG, Salles FF (2005) Three new species of Tricorythopsis

(Ephemeroptera: Leptohyphidae) from Southeastern Brazil. Aquat Insect 27: 235-241.

Dias LG, Salles FF, Ferreira PSF (2008) New species of Tricorythopsis

Traver (Ephemeroptera: Leptohyphidae) from Northern Brazil. Stud Neotrop Fauna Environ 43: 237-241.

Dias LG, Salles FF, Polegatto CM, Silva R, Froehlich CG (2007b) Novos registros de Ephemerelloidea (Insecta: Ephemeroptera) para o estado de São Paulo. Biota Neotrop 7: 37-40.

Domínguez E, Molineri C, Pescador ML, Hubbard MD, Nieto C

(2006) Ephemeroptera of South America. Vol. 2. Sofia-Moscow,

Pensoft, 646p.

Gonçalves IC, Da-Silva ER (2010) Ephemeroptera, Leptohyphidae,

Amanahyphes saguassu Salles and Molineri, 2006: first record

from state of Pará, Brazil. Check List 6: 170-171 .

Instituto Brasileiro de Geografia e Estatística, Ministério do

Planejamento, Orçamento e Gestão, Brasil (2002) Mapa de climas do Brasil. ftp://geoftp.ibge.gov.br/mapas/ tematicos/ mapas_murais/clima.pdf. (Last accessed 12/05/2011).

Instituto Brasileiro de Geografia e Estatística, Ministério do

Planejamento, Orçamento e Gestão, Brasil (2004) Mapa de Biomas do Brasil. ftp://geoftp.ibge.gov.br/mapas/tematicos/ mapas_murais/ biomas.pdf. (Last accessed 12/05/2011).

Leal IR, Silva JMC, Tabarelli M, Lacher TER (2005) Mudando o curso da conservação da biodiversidade na Caatinga do Nordeste do Brasil. Megadiversidade 1: 139-146.

Lima LRC, Salles FF, Pinheiro US, Quinto E (2010) Espécies de

Baetidae (Ephemeroptera) do Sul da Bahia, com descrição de

uma nova espécie e Paracloeodes Day. Neotrop Entomol 39: 725-731

Mariano R, Polegatto C (2011) Checklist de Ephemeroptera do estado de São Paulo, Brasil. Biota Neotrop 11: 291-297.

Molineri C (2001a) Traverhyphes: A new genus of Leptohyphidae for Leptohyphes indicator and related species (Insecta: Ephemeroptera). Spixiana 24: 129-140.

Molineri C (2001b) El género Tricorythopsis (Ephemeroptera:

Leptohyphidae): nuevas combinaciones y descripción de nuevas especies y estádios.Rev Soc Entomol Argent 60: 217-238

Molineri C (2002) Cladistic analysis of the South American species of Tricorythodes (Ephemeroptera: Leptohyphidae) with the description of new species and stages. Aquat Insect 24: 273-308.

Molineri C (2004) Phylogeny of the Allenhyphes-Traverhyphes group (Ephemeroptera: Leptohyphidae), with new subgenera, species and combinations. Tijdschr Entomol 147: 197-220.

Needham JG, Murphy HE (1924) Neotropical Mayflies. Bull Lloyd

Libr, Entomol Ser 24: 1-79.

Rebouças AC (1997) Água na Região Nordeste: desperdício e escassez. Estud Av 11: 127-154.

(7)

Salles FF, Boldrini R, Nascimento JC, Raimundi E, Shimano Y (2010b) Ephemeroptera do Brasil. http://Ephemeroptera.br.googlepages. com/home. (Last accessed 10/12/2010).

Salles FF, Da-Silva ER, Hubbard MD, Serrão JE (2004) As espécies

de Ephemeroptera (Insecta) registradas para o Brasil. Biota Neotrop 4: 1-34.

Salles FF, Molineri C (2006) Amanahyphes saguassu, A new genus and species of Leptohyphidae (Ephemeroptera: Ephemerelloidea) from Northern Brazil. Aquat Insect 28: 1-12.

Salles FF, Nascimento J, Massariol F, Angeli K, Barcelos-Silva P, Rúdio

J, Boldrini R (2010a) First survey of Mayflies (Ephemeroptera,

Insecta) from Espírito Santo State, Southeastern Brazil. Biota Neotrop 10: 293-307.

Silva JMC, Tabarelli M, Fonseca MT, Lins LV (orgs) (2004) Biodiversidade da caatinga: áreas e ações prioritárias para a

conservação. Brasília, Ministério do Meio Ambiente, 382p.